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Anda di halaman 1dari 13

J.M. CUSHING

a,

*, SHEREE LEVARGE

a

, NAKUL CHITNIS

a

and

SHANDELLE M. HENSON

b

a

Department of Mathematics, Program on Applied Mathematics, University of Arizona,

Tucson, AZ 85721, USA;

b

Department of Mathematics, Andrews University, Berrien Springs,

MI 49104, USA

(Received 6 May 2003; In nal form 11 November 2003)

Dedicated to Professor Saber Elaydi on the Occasion of His 60th Birthday

Adifference equation model, called that Leslie/Gower model, played a key historical role in laboratory experiments that

helped establish the competitive exclusion principle in ecology. We show that this model has the same dynamic

scenarios as the famous Lotka/Volterra (differential equation) competition model. It is less well known that some

anomalous results fromthe experiments seemtocontradict the exclusionprinciple andLotka/Volterra dynamics. We give

an example of a competition model that has non-Lotka/Volterra dynamics that are consistent with the anomalous case.

Keywords: Difference equations; Global stability; Competition models; Competitive exclusion principle;

Leslie/Gower model; Lotka/Volterra dynamics

AMS Nos: 39A11; 92D40

INTRODUCTION

A fundamental tenet in theoretical ecology is the competitive exclusion principle.

According to this tenet, two similar species competing for a limited resource cannot coexist;

one of the species will be driven to extinction. This principle is supported by many

mathematical models, the most famous of which is the Lotka/Volterra differential equation

model for two competing species. It is well known that the Lotka/Volterra model allows just

four dynamic scenarios, all of which involve only equilibria as possible asymptotic states.

A coexistence case (in the form of a globally asymptotically stable equilibrium) occurs if the

competition between the species is weak. If, however, the inter-species competition is

sufciently strong, then competitive exclusion occurs (in the form of an equilibrium state

possessing one zero component). The competitive exclusion case has three possible

dynamic scenarios, depending upon relationships among the model coefcients. Two of

these scenarios are symmetric cases that have globally attracting equilibria in which one

species is absent. The third, and nal scenario, has an unstable (saddle) coexistence

Journal of Difference Equations and Applications

ISSN 1023-6198 print/ISSN 1563-5120 online q 2004 Taylor & Francis Ltd

http://www.tandf.co.uk/journals

DOI: 10.1080/10236190410001652739

This research was supported by National Science Foundation grant DMS 9973126.

*Corresponding author. E-mail: cushing@math.arizona.edu

Journal of Difference Equations and Applications,

Vol. 10, No. 1315, November December 2004, pp. 11391151

equilibrium whose stable manifold determines two basins of attraction, one for each of the

competitive exclusion equilibria. In this saddle case, the species that goes extinct is

determined by the initial conditions of both species.

During the 1940s, 50s and 60s, laboratory experiments played a key role in establishing the

competitive exclusion principle in theoretical ecology. One series of laboratory studies, which

today is still cited in text books, was conducted by Park and Mertz using two species of our

beetles (of the genus Tribolium) [3740]. Park and his collaborators, Leslie and Gower, used a

difference equation model in these studies, rather than the Lotka/Volterra differential equations

[26]. Although they did not give a mathematical analysis of their model, they worked under the

assumption that it possesses the same dynamic scenarios as the Lotka/Volterra model.

It is interesting that although Parks experiments were considered a validation of the

competitive exclusion principle, there were some anomalous results. In one experiment

competitive exclusion did not always occur. Whether competitive coexistence occurred

or competitive exclusion occurred (and how) depended on the initial population numbers given

the two species. This result puzzled Park and his collaborators, since this dynamic scenario is not

allowed by Lotka/Volterra-type competition models which guided their thinking [14,27].

A natural question to examine, from a mathematical point of view, is whether or not there exist

any two species competition models that allow for such a multiple attractor, coexistence/exclu-

sion case. Anumber of studies involving competition models have, under certain circumstances,

found results of non-Lotka/Volterra type or that contradicted the competitive exclusion principle

in one way or another [2,58,10,13,20,2224,28,29,3136,41,42]. None of these results,

however, involve the multiple attractor scenario described above.

The Leslie/Gower competition model was studied by Liu and Elaydi, who showed that all

bounded orbits converge to an equilibrium in an eventually monotonic manner (using the

theory of discrete monotone ows) [30]. In this paper we will extend this result by showing

that the Leslie/Gower model has the same dynamic scenarios as the Lotka/Volterra model

(characterizing these cases according to model parameters). We will also investigate a

competition model that does conform not to the Lotka/Volterra cases and that allows for

multiple, coexistence and exclusion, attractors.

THE LESLIE/GOWER MODEL

If b . 1 all solutions of the Beverton/Holt equation

x

t1

b

1

1 c

11

x

t

x

t

with x

0

. 0 tend (monotonically) to the equilibrium x b 21=c

11

: This difference

equation is an appropriate analog of the logistic differential equation [12]. Just as the famous

Lotka/Volterra two species (differential equation) competition model is a modication of the

logistic differential equation, the Leslie/Gower (difference equation) competition model [26]

x

t1

b

1

1

1 c

11

x

t

c

12

y

t

x

t

y

t1

b

2

1

1 c

21

x

t

c

22

y

t

y

t

is a modication of the Beverton/Holt equation. In this model all coefcients are positive and

we can therefore scale x and y so that c

11

c

22

1: Without loss in generality we consider

J.M. CUSHING et al. 1140

the system

x

t1

b

1

1

1 x

t

c

1

y

t

x

t

1

y

t1

b

2

1

1 c

2

x

t

y

t

y

t

2

where b

i

. 0 and c

i

. 0: We denote solutions of this system by x

t

; y

t

; t 0; 1; 2; 3; . . .:

(For some results concerning difference equations dened by rational functions see

Refs. [3,4,25]. The results in these papers do not apply to the Leslie/Gower model, however.

Other papers that deal with discrete competition models include [1619].)

In population dynamic applications we are interested in solutions with non-negative

components x

t

$ 0; y

t

$ 0: Let R

2

z 0; 1 0; 1 and

R

2

z 0; 1 0; 1

f : x; y ! b

1

1

1 x c

1

y

x; b

2

1

1 c

2

x y

y

_ _

takes R

2

R

2

{0} 0; 1: Moreover, all solutions in R

2

f : R

2

!S z 0; b

1

0; b

2

: It follows from Proposition 1 in Ref. [30] that all orbits in

R

2

The map f is also invertible on R

2

; since for x

0

; y

0

[ S in the range of f the equations

b

1

1

1 x c

1

y

x x

0

; b

2

1

1 c

2

x y

y y

0

have the unique solution

x

b

2

21 c

1

D

; y

b

1

21 c

2

D

where

b

1

z

b

1

x

0

. 1; b

2

z

b

2

y

0

. 1; D z b

1

21b

2

21 2c

1

c

2

(the range of f is dened by the inequality D . 0). The formulas for the pre-images x and y

show the inverse f

21

continuous.

Lemma 1 The map f : R

2

The points E

0

: 0; 0; E

1

: b

1

21; 0; E

2

: 0; b

2

21 are xed points of the map f (i.e.

are equilibria of the Leslie/Gower model (1) and (2)). These are exclusion equilibria.

The set of points whose x-coordinate is held xed by the map f is the line x c

1

y

b

1

21: If this line intersects

R

2

(i.e. if b

1

. 1), we denote the resulting line segment by L

1

.

Similarly, if b

2

. 1; the points on the line segment L

2

from the line c

2

x y b

2

21 lying in

R

2

2

1

. 1 the map

f takes a point x; y [ R

2

1

to a point with smaller (larger)

x-coordinate. If b

2

. 1 the map f takes a point x; y [ R

2

2

to a point

with smaller (larger) y-coordinate.

The only other xed point of f is

E

3

:

b

2

21

c

1

c

2

21

c

1

2

b

1

21

b

2

21

_ _

;

b

1

21

c

1

c

2

21

c

2

2

b

2

21

b

1

21

_ _ _ _

COMPETITION MODELS AND EXCLUSION PRINCIPLE 1141

This equilibrium lies in R

2

if and only if b

1

. 1; b

2

. 1 and L

1

and L

2

intersect in R

2

:

This is a coexistence equilibrium.

An equilibrium is globally asymptotically stable on R

2

(or

R

2

) if it is locally

asymptotically stable (on R

2

) and if x

0

; y

0

[ R

2

(or

R

2

) implies that x

t

; y

t

tends to the

equilibrium as t !1:

The Jacobian of Eqs. (1) and (2) is

J

2b

1

1

1xc

1

y

2

x b

1

1

1xc

1

y

2b

1

c

1

1

1xc

1

y

2

x

2b

2

c

2

1

1c

2

xy

2

y 2b

2

1

1c

2

xy

2

y b

2

1

1c

2

xy

_

_

_

_

_

_

The Jacobians evaluated at E

0

, E

1

and E

2

are

J

0

b

1

0

0 b

2

_ _

; J

1

1

b

1

1

b

1

1 2b

1

c

1

0

b

2

1b

1

21c

2

_

_

_

_

; J

2

b

1

1b

2

21c

1

0

1

b

2

1 2b

2

c

2

1

b

2

_

_

_

_

respectively. Their eigenvalues appear along the diagonals.

Lemma 2

(a) If b

1

, 1; b

2

, 1 then E

0

is globally asymptotically stable on R

2

:

(b) If b

1

. 1; b

2

, 1 then E

1

is globally asymptotically stable on

R

2

:

(c) If b

1

, 1; b

2

. 1 then E

2

is globally asymptotically stable on

R

2

:

Proof

(a) The eigenvalues of J

0

are less than 1 and E

0

is locally asymptotically stable. Since E

0

is

the only equilibrium in R

2

it follows from the Liu and Elaydis theorem that all solutions

in R

2

converge to E

0

. This can also be seen, more directly, from the inequalities

0 # x

t1

, b

1

x

t

3

0 # y

t1

, b

2

y

t

4

and an induction argument.

(b) The only equilibria in R

2

are E

0

and E

1

. From J

0

we see that E

0

is a saddle. Since the

coordinate axes are invariant, the stable manifold is the y-axis (since by Eq. (4) y

t

!0).

From the stable manifold and Hartman/Grobman theorems for maps [15,21], it follows

that no solution in

R

2

can approach E

0

. By Liu and Elaydis theorem all solutions must

therefore approach E

1

. The eigenvalues of J

1

are less than 1 and E

1

is locally

asymptotically stable.

(c) This symmetric case is proved in manner analogous to case (b). A

From now on we assume b

1

. 1; b

2

. 1: These inequalities imply E

1

and E

2

lie on the

positive x- and y-axis, respectively. We distinguish four cases depending on the orientation

of the line segments L

1

and L

2

as shown in Figure 1. The inequalities satised by the

coefcients that correspond to these cases can be easily read from the relative positions of

the intercepts of L

1

and L

2

in Figure 1. In particular, the following inequalities characterize

J.M. CUSHING et al. 1142

Cases B and C:

Case B : c

1

b

2

21 , b

1

21; c

2

b

1

21 , b

2

21

Case C : c

1

b

2

21 . b

1

21; c

2

b

1

21 . b

2

21:

5

From these inequalities we nd that

c

1

c

2

, 1 holds in Case B

c

1

c

2

. 1 holds in Case C:

6

Lemma 3 Assume b

1

. 1 and b

2

. 1: The equilibrium E

0

is a repellor. For i 1; 2 the

equilibrium E

i

is locally asymptotically stable in Cases A

i

and C, but is a saddle in Cases A

j

j i and B. In the latter two cases, the stable manifold of the saddle E

i

is the positive

coordinate axis on which it lies. Equilibrium E

3

is locally asymptotically stable in Case B

and is a saddle in Case C.

Proof The eigenvalues of J

0

are b

1

. 1; b

2

. 1 and E

0

is therefore a repellor.

The eigenvalue 1/b

1

of J

1

is less than one. Solutions starting on the positive x-axis

satisfy the Bervton/Holt equation (with b

1

. 1) and therefore approach E

1

. The second

eigenvalue

b

2

1 c

2

b

1

21

is less than one in Cases A

1

and C and greater than one in Cases A

2

and B. A similar

calculation shows the result for E

2

.

FIGURE 1 The four possible orientations of L

1

and L

2

.

COMPETITION MODELS AND EXCLUSION PRINCIPLE 1143

Finally, we consider the equilibrium E

3

, at which the Jacobian is

J

3

c

1

c

2

b

1

2c

1

b

2

c

1

21

b

1

c

1

c

2

21

c

1

b

1

2c

1

b

2

c

1

21

b

1

c

1

c

2

21

c

2

2c

2

b

1

b

2

c

2

21

b

2

c

1

c

2

21

2c

2

b

1

c

2

c

1

b

2

c

2

21

b

2

c

1

c

2

21

_

_

_

_

_

_

_

_

_

_

:

This equilibrium is locally asymptotically stable if the Jury conditions

jtr J

3

j , 1 det J

3

, 2

hold, where det J

3

is the determinant and tr J

3

is the trace of J

3

[1]. If at least one of these

inequalities is reversed, then the equilibrium is unstable. The inequalities are equivalent to

the following three inequalities

(a) 1 det J

3

, 2

(b) 21 2det J

3

, tr J

3

(c) tr J

3

, 1 det J

3

.

A calculation shows

det J

3

c

2

c

1

21b

1

21 c

1

c

2

21b

2

21 c

1

c

2

21

b

1

b

2

c

1

c

2

21

tr J

3

2b

2

b

1

c

1

c

2

2b

2

c

1

b

2

2c

1

b

2

2

2b

1

c

2

b

1

2c

2

b

2

1

b

1

b

2

c

1

c

2

21

:

Using these formulas, we nd that inequality (a) is equivalent to

b

1

21b

2

21

c

1

b

2

21 2b

1

21

c

1

c

2

21

c

2

b

1

21 2b

2

21

c

1

c

2

21

. 0:

By Eqs. (5) and (6) this inequality holds. Inequality (b) is equivalent to

b

1

1b

2

1 c

2

b

1

1

c

1

b

2

21 2b

1

21

c

1

c

2

21

c

1

b

2

1

c

2

b

1

21 2b

2

21

c

1

c

2

21

.0

which Eqs. (5) and (6) show also holds true. Finally, we consider inequality (c), which it turns

out is equivalent to

c

1

b

2

21 2b

1

21c

2

b

1

21 2b

2

21

c

1

c

2

21

,0:

From Eqs. (5) and (6), we see that this inequality holds in Case B and, as a result, E

3

is locally

asymptotically stable. In Case C, however, the reverse inequality holds and E

3

is unstable.

To see, in the latter case, that E

3

is in fact a saddle we note that the characteristic quadratic

polynomial of J

3

, namely,

pl l

2

2tr J

3

l det J

3

J.M. CUSHING et al. 1144

satises p1 1 2tr J

3

det J

3

, 0 (because the inequality in (c) is reversed in Case C),

p21 1 tr J

3

det J

3

. 0 (because (b) holds), and p1 1: Thus, p(l) has a

positive root l . 1 and a root between 21 and 1. A

Theorem 4 Consider the Leslie/Gower competition model (1) and (2) with b

i

. 1:

Each component x

t

and y

t

of a solution in

R

2

is eventually monotonic.

(a) In Case A

i

, the equilibrium E

i

is globally asymptotically stable on

R

2

:

(b) In Case B, the equilibrium E

3

is globally asymptotically stable on

R

2

:

(c) In Case C, for solution in

R

2

E

1

or E

2

or to the saddle equilibrium E

3

as t !1:

Proof Every solution in

R

2

equilibria E

i

by Liu and Erlaydis theorem.

(a) The two Cases A

1

and A

2

are symmetric and we consider Case A

1

only. Observe that the

(closed) triangular region E

0

E

1

P is forward invariant under the map f. This follows

from two facts: f is one-to-one and bicontinuous (Lemma 1) and the boundary of E

0

E

1

P

maps into E

0

E

1

P (see Fig. 1). Thus, if a solution starts in the triangle E

0

E

1

P is remains

there and as a result the component x

t

is increasing. It follows that the solution

approaches E

1

. If, on the other hand, x

0

; y

0

[

R

2

0

E

1

P there

are two options: either the solution sequence x

t

; y

t

enters the triangle in a nite number

of steps, in which case it will remain there and converge to E

1

(as we just proved), or it

remains outside the triangle for all t. In the latter case, the solution can only converge to

E

1

. In summary, x

0

; y

0

[

R

2

implies x

t

; y

t

tends E

1

as t !1: This, together with

Lemma 3, shows E

1

is globally asymptotically stable on

R

2

:

(b) By Lemma 3, E

3

is locally asymptotically stable. The discrete stable manifold and

Hartman/Grobman theorems, together with Lemma 3, imply that no orbit in

R

2

can

approach E

0

, E

1

or E

2

. It follows from Liu and Elaydis theorem that all orbits in

R

2

approach E

3

.

(c) By Lemma 3, E

0

is a repellor and cannot be approached by a solution in

R

2

: A

For case (c) in Theorem 4 we do not have a complete characterization of the basins of

attraction for each equilibrium. From the stable manifold theorem for maps [15,21] we know

the local stable manifold of E

3

is one dimensional (since by Lemma 2 the equilibrium E

3

is a

hyperbolic saddle in Case C), but we do not know if this is globally true in R

2

: We can show

that the basin of attraction of E

1

contains the interior of the closed triangle QE

1

E

3

. To see this

notice that f maps the boundary of the triangle into the triangle: f : QE

1

E

3

!QE

1

E

3

(see Fig. 1). Since f is oneone and bicontinuous, it follows that f : QE

1

E

3

!QE

1

E

3

(i.e. QE

1

E

3

is forward invariant). Solutions starting in (or eventually entering) the interior of

QE

1

E

3

must accordingly approach the equilibrium E

1

. Similarly, one can show that the basin

of attraction of E

2

contains the interior of the triangle PE

2

E

3

.

The four dynamic cases in Theorem 4 (and Fig. 1) are the same as those in the

Lotka/Volterra model and have the same ecological interpretation. Coexistence (Case B)

occurs if interspecic competition is weak in the sense that the coefcients c

i

are small:

c

1

,

b

1

21

b

2

21

; c

2

,

b

2

21

b

1

21

:

COMPETITION MODELS AND EXCLUSION PRINCIPLE 1145

If interspecic competition is too large (one or both inequalities are reversed), then

competitive exclusion occurs. This is the theoretical foundation for the classical competitive

exclusion principle. It is because of this principle that the anomolous case observed in some

of Parks experiments were puzzling. In the next section, we will study a competition model

which does not support this exclusion principlea model which in fact predicts (non-

equilibrium) coexistence when the competition coefcients are increased.

A JUVENILE/ADULT RICKER MODEL

Since the Leslie/Gower competition model (1) and (2) has only the Lotka/Volterra dynamics

described in Theorem 4, this model offers no explanation of the multiple attractor case

(with initial condition dependent coexistence or exclusion) observed by Park. A discrete

competition model that does provide an explanation is the so-called LPA model [1012].

This six dimensional model is based on three life cycle stages for each species and was

derived explicitly to study the dynamics of our beetle species. See Ref. [14] for an account

of the LPA competition model and some of its non-Lotka/Volterra dynamics that contradict

the classical competitive exclusion principle.

A mathematical exploration of competition models that possess multiple attractor cases

with initial condition dependent coexistence or exclusion might start with the Leslie/Gower

model and try to determine what kinds of structural changes in the model will result in

non-Lotka/Volterra dynamics and contradictions to the competitive exclusion principle.

For example, if exponential non-linearities replace the rational function non-linearities of the

Leslie/Gower model, one obtains the Ricker competition model

x

t1

b

1

x

t

exp2c

11

x

t

2c

12

y

t

y

t1

b

2

y

t

exp2c

21

x

t

2c

22

y

t

(exponential non-linearities also appear in the LPA model). Numerical investigations of this

model have uncovered certain kinds of non-Lotka/Volterra dynamics [10], but not a case of

multiple attractors with both coexistence and exclusion. Whether or not such multiple

attractor cases can occur in this system remains an interesting open question.

From a mathematical point of view life cycle stages (such as appear in the LPA model)

introduce time delays. A modication of the Ricker competition model, in which individuals

from one of the two species are characterized by their reproductive maturity, is described by

the difference equations

J

t1

b

1

A

t

exp2c

11

A

t

2c

12

y

t

A

t1

1 2mJ

t

y

t1

b

2

y

t

exp2c

21

J

t

2c

22

y

t

:

7

Here J

t

and A

t

are the numbers of juveniles and adults at time t of the species x. Species y in

this model remains unstructured. The parameter m 0 # m , 1 is the juvenile mortality rate.

In this paper, we will not attempt a thorough analysis of the juvenile/adult Ricker

competition model (7). We will only show that under certain conditions the model possesses

three attractors, two exclusion (equilibrium) attractors and one (non-equilibrium)

coexistence attractor.

J.M. CUSHING et al. 1146

The exclusion equilibria for Eq. (7) are

E

0

: J; A; y 0; 0; 0

E

1

: J; A; y

ln n

c

11

1 2m

;

ln n

c

11

; 0

_ _

E

2

: J; A; y 0; 0;

ln b

2

c

22

_ _

:

Here we have dened n z b

1

1 2m [9]. We assume n . 1; b

2

. 1 so that these equilibria

are non-negative. The Jacobian at E

0

0 b

1

0

2m 1 0 0

0 0 b

2

_

_

_

_

_

_

_

_

has eigenvalues l b

2

and ^

n

p

: The Jacobian at E

1

0

12ln n

n

b

1

2

b

1

c

12

ln n

c

11

n

1 2m 0 0

0 0 b

2

exp 2

c

21

ln n

c

11

12m

_ _

_

_

_

_

_

_

_

_

_

_

has eigenvalues l ^

1 2ln n

p

and b

2

exp(2c

21

ln n/c

11

(1 2 m)). The Jacobian at E

2

0 b

1

e

2

c

12

c

22

ln b

2

0

1 2m 0 0

2

c

21

c

22

ln b

2

0 1 2ln b

2

_

_

_

_

_

_

_

_

_

_

has eigenvalues l ^

n

p

and 1 2 ln b

2

.

Theorem 5 Assume n b

1

1 2m . 1; b

2

. 1 in the juvenile/adult Ricker competition

model (7).

(a) E

0

is a repellor.

(b) E

1

is locally asymptotically stable if

1 , n , e

2

; c

11

1 2m

ln b

2

ln n

, c

21

:

(c) E

2

is locally asymptotically stable if

1 , b

2

, e

2

; c

22

ln n

ln b

2

, c

12

:

The ecological interpretation of the parameter inequalities in Theorem 5(b) and (c) is that

interspecic competition is sufciently strong (as measured by c

12

and c

21

), relative to

intraspecic competition (measured by c

11

and c

22

). This theorem suggests the

Lotka/Volterra or Leslie/Gower saddle case of competitive exclusion (Case C in Fig. 1).

However, it is possible that all the inequalities in Theorem 5 hold and there also exists

COMPETITION MODELS AND EXCLUSION PRINCIPLE 1147

a coexistence attractor. Figure 2 shows an example. In that gure one initial condition

approaches a coexistence two-cycle, while other initial conditions lead to the extinction

equilibria E

1

and E

2

.

It is possible to prove the existence of coexistence two-cycles of the type observed in Figure 2.

Two-cycles are xed points of the composite map dened by the model equations (7)

b

1

1 2mJ exp2c

11

1 2mJ 2b

2

c

12

ye

2c

21

J2c

22

y

J

b

1

1 2mAe

2c

11

A2c

12

y

A

b

2

2

ye

2c

21

J2c

22

y

exp2b

1

c

21

Ae

2c

11

A2c

12

y

2b

2

c

22

ye

2c

21

J2c

22

y

y:

8

Although it is not shown in Fig. 2, the coexistence two-cycle in that example is

synchronous, i.e. the two points of the cycle are of the form

0; A; y

1

; J; 0; y

2

9

with J . 0; A . 0; and y

i

. 0: Synchronous means that the juvenile and adult populations

never appear together at the same point in time. The two-cycle equations (8) with J 0

reduce to the two equations

b

1

1 2me

2c

11

A2c

12

y

1

b

2

2

e

2c

21

J2c

22

y

exp2b

1

c

21

Ae

2c

11

A2c

12

y

2b

2

c

22

ye

2c

21

J2c

22

y

1

FIGURE 2 Three different initial conditions lead to solutions of the juvenile/adult Ricker competition model (7)

that approach three different attractors. (a) The initial condition J

0

; A

0

; y

0

10; 10; 10 results in an orbit that

approaches an equilibrium in which y 0; i.e. this initial condition leads to the extinction of species y. (b) The initial

condition J

0

; A

0

; y

0

5; 5; 10 results in an orbit that approaches an equilibrium in which J A 0; i.e. this

initial condition leads to the extinction of species x. (c) The initial condition J

0

; A

0

; y

0

10; 10; 15 results in an

orbit that approaches a positive two-cycle, i.e. the initial condition leads to the (non-equilibrium) coexistence of

both species. The coefcients in these examples are b

1

b

2

5; c

11

c

22

0:1; c

12

0:11; c

21

0:12; and

m 0:2:

J.M. CUSHING et al. 1148

for A and y. Using the rst equation in the second we can simplify the second equation

b

1

1 2me

2c

11

A2c

12

y

1

b

2

2

exp 2c

22

y 2c

21

1

1 2m

A 2b

2

c

22

ye

2c

22

y

_ _

1:

Solving the rst equation for

A

1

c

11

ln n 2c

12

y 10

substituting this result into the second equation and simplifying, we arrive at the single

equation

b

2

c

22

ye

2c

22

y

c

12

c

21

1 2mc

11

2c

22

_ _

y 2 ln b

2

2

c

21

1 2mc

11

ln b

1

1 2m

_ _

11

for y. A solution y y

1

. 0; together with Eq. (10), yields the rst point in a two-cycle (9).

(If A . 0 then this point is not an equilibrium.) The Eq. (11) can be analyzed geometrically

by investigating the graphs of both sides of the equation for intersection points y . 0: The left

hand side is a positive, one humped graph passing through y 0 and having y 0 as an

asymptote. The right hand side is a straight line whose slope is positive under the

assumptions in Theorem 5(b) and (c). If the y-intercept of the straight line is positive, then

either there are two intersection points of these graphs, no intersection point at all, or a

tangency case of one intersection point. As parameters change and we pass from one to the

other case, a bifurcation occurs at the point of tangency. (We speculate that this is a saddle

node bifurcation with another two-cycle which, because it is not a synchronous cycle, is not

found by the equations above.) See Figure 3 for an illustration. Note that the coexistence two-

cycle is created in this example by increasing the interspecic competition coefcient c

21

!

FIGURE 3 Roots y . 0 of Eq. (9) determine synchronous two-cycles for the juvenile/adult Ricker competition

model (7). The dashed line graph is that of the left hand side of this equation for the parameter values in Fig. 2.

The straight line graphs are the right hand side of the equation for three selected values of the interspecic

competition coefcient c

21

. For the smaller value of c

21

there is no intersection and there exists no synchronous two-

cycle. For the larger value of c

21

, two intersection points exist and give rise to a synchronous two-cycle.

COMPETITION MODELS AND EXCLUSION PRINCIPLE 1149

To establish the triple attractor case of interest (and observed in Fig. 2) we need to prove

the stability of the two-cycle under the conditions of Theorem 5(b) and (c). The two-cycle is

locally asymptotically stable if the eigenvalues of the Jacobian of the composite map are less

than one in magnitude. We can calculate this Jacobian by multiplying the Jacobians of the (7)

evaluated at the two points of the two-cycle. In general, this calculation is intractable, but we

can carry it out for the example in Figure 2. For the parameter values in that example, Eq. (11)

has the two solutions y

1

4:4758 and y

2

14:304 which produce the two-cycle points

(see Eq. (9))

0; 8:9396; 4:4758; 11:1174; 0; 14:304:

The Jacobians of Eq. (7) at these two points, when multiplied, yield the matrix

0:76624 0 0:16554

0 0:82932 0

20:94822 21:7794 20:23776

_

_

_

_

_

_

_

_

whose eigenvalues l 24:4039 10

22

; 0.57252, and 0.82932 are less than one in

magnitude. The two-cycle is therefore locally asymptotically stable. Since the inequalities

in Theorem 5(b) and (c) are satised by the parameters in this example the two exclusion

equilibria

E

1

: J; A; y 20:118; 16:094; 0

E

2

: J; A; y 0; 0; 16:094

are also locally asymptotically stable.

The juvenile/adult Ricker competition model shows that strong non-linearities and a time

delay can produce a multiple attractor, coexistence/exclusion, scenario. According to our

analysis above this scenario arises from a bifurcation that gives rise to a two-cycle

coexistence state as interspecic competition is increased, which from the point of view of

classical competition theory is unexpected. This same bifurcation scenario occurs in the LPA

model [14]. Thus, we have two examples of competition models that provide dynamics

which contradict classical competition theory in signicant ways. An interesting open

mathematical problem is to determine the types of competition models that possess these

kinds of multiple attractors.

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COMPETITION MODELS AND EXCLUSION PRINCIPLE 1151

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